The Lichens of British Columbia, Illustrated Keys, Part 1 - Foliose and Squamulose Species [8] 0772621942

Table of contents :
Cover
TABLE OF CONTENTS
ACKNOWLEDGEMENTS
INTRODUCTION
Interpreting the Species Accounts
Understanding Biogeoclimatic Zonation
Identifying Lichens
Making Use of Lichen Chemistry
A Note on Common Names
KEYS TO GENERA OF FOLIOSE AND SQUAMULOSE LICHENS
Making Use of the Keys
KEY A: LICHEN GROWTH FORMS
KEY B: NONSTRATIFIED (OR APPARENTLY NONSTRATIFIED) FOLIOSE AND SQUAMULOSE LICHEN GENERA OF BRITISH COLUMBIA
“Gel Lichens”
KEY C: STRATIFIED SQUAMULOSE LICHEN GENERA OF BRITISH COLUMBIA
“Scale Lichens”
KEY D: STRATIFIED FOLIOSE LICHEN GENERA OF BRITISH COLUMBIA
KEYS TO SPECIES OF FOLIOSE AND SQUAMULOSE LICHENS, BY GENUS
AGONIMIA
AHTIANA
ALLANTOPARMELIA
ANAPTYCHIA
ARCTOPARMELIA
ASAHINEA
BRODOA
CANDELARIA
CATAPYRENIUM
CAVERNULARIA
CETRARIA
CETRELIA
COLLEMA
DERMATOCARPON
ENDOCARPON
ERIODERMA
ESSLINGERIANA
FLAVOPUNCTELIA
GONOHYMENIA
HEPPIA
HETERODERMIA
HYDROTHYRIA
HYPOCENOMYCE
HYPOGYMNIA
HYPOTRACHYNA
IMSHAUGIA
KOERBERIA
LASALLIA
LEIODERMA
LEPTOCHIDIUM
LEPTOGIUM
LOBARIA
MASSALONGIA
MELANELIA
MENEGAZZIA
NEOFUSCELIA
NEPHROMA
NORMANDINA
PANNARIA
PARMELIA
PARMELIELLA
PARMOTREMA
PELTIGERA
PELTULA
PHAEOPHYSCIA
PHYLLISCUM
PHYSCIA
PHYSCONIA
PLACYNTHIUM
PLATISMATIA
PSEUDOCYPHELLARIA
PSORA
PSOROMA
PUNCTELIA
RHIZOPLACA
SOLORINA
STICTA
UMBILICARIA
VESTERGRENOPSIS
VULPICIDA
WAYNEA
XANTHOPARMELIA
XANTHORIA
APPENDIX 1. Distribution maps of rare and infrequent foliose and squamulose lichens
APPENDIX 2. Excluded species
GLOSSARY AND ABBREVIATIONS
REFERENCES
INDEX
Figures
FIGURE 1. First- and second-order lichen floristic studies in British Columbia to 1992.
FIGURE 2. “Life zones” of British Columbia.
FIGURE 3. Biogeoclimatic zones of British Columia
FIGURE 4. Thallus stratified/heteromerous
FIGURE 5. Thallus nonstratified/homoiomerous
FIGURE 6. Organs of attachment
FIGURE 7. Lichen growth forms:
FIGURE 8. Surface details
FIGURE 9. Reproductive structures
Tables
TABLE 1. Distributional units and their definition
TABLE 2. Summary information on the biogeoclimatic zones of British Columbia

Citation preview

Special Report Series

8

THE LICHENS OF BRITISH COLUMBIA Illustrated Keys Part 1 — Foliose and Squamulose Species

Ministry of Forests Research Program

1 9 9 4 ISSN 0843-6452

The Lichens of British Columbia Illustrated Keys Part 1 — Foliose and Squamulose Species

by

Trevor Goward, Bruce McCune, and Del Meidinger (Illustrations by Trevor Goward)

Ministry of Forests Research Program

1 9 9 4

AUTHORS AND AFFILIATIONS This report has been prepared for the Research Program, Ministry of Forests by: Trevor Goward Herbarium, Department of Botany University of British Columbia Vancouver, B.C. V6T 1Z4 Mailing Address: Edgewood Blue, Box 131 Clearwater, B.C. V0E 1N0 Bruce McCune Department of Botany and Plant Pathology Cordley Hall 2082 Oregon State University Corvallis, Oreg. 97331-2902 Del Meidinger B.C. Ministry of Forests 31 Bastion Square Victoria, B.C. V8W 3E7

Canadian Cataloguing in Publication Data Goward, Trevor. The lichens of British Columbia, illustrated keys. Part 1, Foliose and Squamulose species. (Special report series, ISSN 0843-6452 ; 8) Includes bibliographical references: p. 181 ISBN 0-7726-2194-2 1. Lichens - British Columbia - Identification. 2. Lichens - British Columbia - Geographic distribution. I. McCune, Bruce. II. Meidinger, Dellis Vern, 1953. III. British Columbia. Ministry of Forests. Research Branch. IV. Title. V. Series: Special report series (British Columbia. Ministry of Forests) ; 8. QK587.7.B7G68 1994

589.1'09711

C94-960252-3

© 1994 Province of British Columbia. 1996 Reprinted with corrections. Published by the Research Program Ministry of Forests 31 Bastion Square Victoria, B.C. V8W 3E7 Copies of this and other Ministry of Forests titles are available from Crown Publications Inc., 546 Yates Street, Victoria, B.C. V8W 1K8.

ii

ACKNOWLEDGEMENTS We are grateful to several people for assistance with this project. Teuvo Ahti and Irwin Brodo kindly reviewed the entire text for accuracy, while several other lichenologists commented on individual genus accounts: Othmar Breuss (Catapyrenium), Gunnar Degelius (Collema), Theodore Esslinger (Melanelia, Neofuscelia, Physcia, Physconia and Phaeophyscia), Per Magnus Jørgensen (Leptogium, Pannaria and Parmeliella), Bruce Ryan (Rhizoplaca) and Einar Timdal (Hypocenomyce and Psora). Chiska Derr and students of the 1993 lichen class at Oregon State University also provided comments on an earlier draft of the keys. Teuvo Ahti, Brian Coppins, Theodore Esslinger, Bernard Goffinet, Thomas Nash, John Thomson, Einar Timdal, Tor Tønsberg and William Weber provided taxonomic opinion on various critical specimens. In addition: Olivia Lee, of the University of British Columbia, packaged, labelled, sorted and shipped thousands of lichen specimens to the senior author, and also cheerfully responded to his occasional requests for lichen literature; Helen Knight spent many hours helping to produce the dot maps; Irwin Brodo and Pak Yau Wong were gracious hosts at the National Museum of Natural Sciences (CANL); Robert Bringhurst, Irwin Brodo, Karen McKeown, Richard Hebda and Roger Rosentreter commented on earlier drafts of proposed common names; Linda Geiser (U.S. Forest Service) kindly made available various unpublished dot maps of the lichens of southeast Alaska; and Yorke Edwards and Andy MacKinnon provided funding through the Royal British Columbia Museum and B.C. Ministry of Forests, respectively, for the drawings that accompany the text. For their help in processing and proofreading the text, as well as bringing the maps and figures to camera-ready, we thank Heather Strongitharm, Susan Bannerman, Malcolm Martin, Hannah Nadel, Merton Palmer, Steve Smith, Paul Nystedt and Dave G. Butcher. Finally, this manual is fondly dedicated to Wilfred Schofield, of the University of British Columbia, whose continued support and friendship have directly and indirectly made it possible. Thanks Wilf!

iii

TABLE OF CONTENTS ACKNOWLEDGEMENTS ........................................................................................................................................

iii

INTRODUCTION ...................................................................................................................................................... Interpreting the Species Accounts ........................................................................................................... Understanding Biogeoclimatic Zonation .................................................................................................. Identifying Lichens ................................................................................................................................... Making Use of Lichen Chemistry ............................................................................................................. A Note on Common Names .....................................................................................................................

1 2 5 10 13 14

KEYS TO GENERA OF FOLIOSE AND SQUAMULOSE LICHENS ....................................................................... Making Use of the Keys ........................................................................................................................... Key A: Lichen Growth Forms .................................................................................................................. Key B: Nonstratified Foliose and Squamulose Lichen Genera of British Columbia ................................ Key C: Stratified Squamulose Lichen Genera of British Columbia ......................................................... Key D: Stratified Foliose Lichen Genera of British Columbia ..................................................................

15 15 16 17 18 22

KEYS TO SPECIES OF FOLIOSE AND SQUAMULOSE LICHENS, BY GENUS ..................................................

30

APPENDIX 1. Distribution maps of rare and infrequent foliose and squamulose lichens in British Columbia ....... 141 APPENDIX 2. Excluded species ............................................................................................................................ 163 GLOSSARY AND ABBREVIATIONS ....................................................................................................................... 165 REFERENCES ......................................................................................................................................................... 169 INDEX .................................................................................................................................................................... 175

TABLES 1 Distributional units and their definition .............................................................................................................

3

2 Summary information on the biogeoclimatic zones of British Columbia ..........................................................

5

FIGURES 1 First- and second-order lichen floristic studies in British Columbia to 1992 .....................................................

1

2 “Life zones” of British Columbia .......................................................................................................................

4

3 Biogeoclimatic zones of British Columbia ........................................................................................................

9

4 Thallus: stratified/heteromerous .......................................................................................................................

10

5 Thallus: nonstratified/homoiomerous ..............................................................................................................

10

6 Organs of attachment .......................................................................................................................................

11

7 Lichen growth forms .........................................................................................................................................

12

8 Surface details .................................................................................................................................................

12

9 Reproductive structures ...................................................................................................................................

13

v

INTRODUCTION Approximately 1100 species of lichens have been reported to occur in British Columbia (B.C.). Although this figure may appear impressive, lichens are among the most poorly documented elements of the province’s macroscopic flora. Judging from the rate at which new species are being added to the lichen flora, it seems likely that hundreds of additional lichens await discovery in this province. Moreover, our understanding of the frequency status of the vast majority of species remains dolefully incomplete. To date, comprehensive lichen studies have been conducted in only two regions of the province: the Queen Charlotte Islands and southeast Vancouver Island. The macrolichen flora of Wells Gray Park is also reasonably well documented. Most of the remainder of the province has received scant attention. Important collections have been made in the regions indicated in Figure 1, but most of these studies are unpublished and the specimens are now scattered in various herbaria. A major impediment to the study of lichens in British Columbia is the lack of comprehensive keys to the species. This manual helps to correct this situation by providing illustrated keys to all “leaf” and “scale” (foliose and squamulose) lichens known to occur in the province. In total, 327 species are included, while 19 taxa are excluded from earlier accounts of the flora. Future volumes in this series will provide keys to the fruticose and crustose species. This manual has two primary objectives. The first is to stimulate lichenological research by making the province’s lichens accessible to as broad an audience as possible. To this end, the keys are tailored primarily to the needs and resources of ecologists, biologists, naturalists, teachers and other non-lichenologists wishing to identify lichens. These users can be assumed: (1) to lack access to thin-layer chromatography (TLC) facilities, as well as to various chemical reagents, ultraviolet lamps, light microscopes and/or other apparatus of detailed lichen identification; (2) to be reluctant

6

10

Figure 1 First- and Second-order Lichen Floristic Studies in British Columbia to 1992*

5 9

10 6

FIRST ORDER 1. Brodo (unpublished) 2. Goward, Ahti (1992) 3. Noble (1982) (Bird, Bird 1973) (Ryan 1991)

Scale 50

8

0

50

100 150 200

kilometres

6

1

SECOND ORDER 4. Benton, Brodo, Richardson (1977) 5. Brodo (unpublished) 6. Goward (unpublished) 7. Goward, Schofield (1983) 8. Ohlsson (unpublished) 9. Otto (unpublished) 10.Thomson, Ahti (unpublished) 130o

* Source: Goward 1993

2 8

9

6 9 3 7

6 4

120o

FIGURE 1. First- and second-order lichen floristic studies in British Columbia to 1992.

INTRODUCTION

to handle unnecessary technical jargon; and (3) to be unfamiliar with basic lichen taxonomy. Based on these assumptions, the keys in this manual: • emphasize morphological characters over chemical and spore characters; • incorporate technical terms only where necessary1; and • give more or less equal weight to phylogenetic relatedness and morphological similarity. Accompanying the keys are approximately 350 line drawings. These are intended to convey species concepts based on typical material. In most cases, they illustrate only those portions of a thallus that bear the characters expressed in the adjacent key. Illustrations of whole lichens may be found in Hale (1979), MacKinnon et al. (1992), Pojar and MacKinnon (1994), Thomson (1984) and Vitt et al. (1988). The manual’s second objective is to briefly summarize the ecology, distribution and frequency status of the province’s foliose and squamulose lichens. Until the status and ecological requirements of lichens are understood, resource managers will have little hope of intelligently managing for lichen diversity. For this reason, distribution maps are provided for species considered vulnerable to logging, grazing, urban development and other forms of human activity. These maps are based primarily on specimens housed at the University of British Columbia, in Vancouver, and the National Museum of Natural Sciences, in Ottawa, though reliable literature reports are also incorporated in some cases. It is beyond the scope of this manual to provide a comprehensive summary of the biology of lichens (see instead: Hale 1983; Hawksworth and Hill 1984; Lawrey 1984). Effective identification does, however, require a basic understanding of lichen morphology and chemistry. The reader is therefore urged to consult the remainder of the Introduction before attempting to use the keys. This manual represents a first attempt to provide comprehensive keys to the province’s foliose and squamulose lichens. The keys have been tested by friends, colleagues and students, but numerous errors and oversights doubtless remain. The user is invited to bring these to the authors’ attention for the benefit of future users.2 Interpreting the Species Accounts Lichens may be arranged into as many as seven different lichen growth forms (see “Identifying Lichens,” page 10). Because these are units of convenience rather than biological units, it is not surprising that some lichen genera embrace more than one growth form. The following accounts incorporate all genera known to occur in British Columbia in which a majority of the species can be described as foliose or squamulose. In a few instances, foliose and squamulose species from other essentially crustose or fruticose genera are also included in the keys. These appear in parentheses (...) and are not discussed in the species accounts. Species appearing in square brackets [...] have not been reliably recorded in the province, but are expected to occur here. These may or may not be discussed in the text. The body of the manual consists of genus and species accounts. These accounts are arranged alphabetically first by genus and then by species within each genus. Each genus account provides: • a common name; • a short description of the genus, with diagnostic characters placed in bold italic type; • pertinent references; • notes on the derivation of the common name; and • notes on global status and distribution, taxonomy, chemistry and/or similar genera.

1

Technical terms are discussed in “Identifying Lichens” (page 10) and appear in boldface at first mention. Additional terms are defined in the keys, as well as in the Glossary (page 165).

2

Please direct comments to Trevor Goward, Edgewood Blue, Box 131, Clearwater, B.C., V0E 1N0

2

INTRODUCTION

The species accounts are organized in the following manner: Species and Author Citation (Synonym): Except in cases of recent taxonomic or nomenclatural revision, species names and author citations follow Egan (1987, 1989, 1990, 1991). Only synonyms in recent and/or widespread use are given. Distribution Maps: Distribution maps are provided (in Appendix 1) for species judged to be of rare or localized occurrence in the province. Map numbers appear to the right of the species names. Common Names: Common names are adopted, adapted or introduced for all lichen species included in this manual. Names given in parentheses (...) have been used by previous authors, but are not accepted here. See also “A Note on Common Names,” page 14. Habitat: Habitat descriptions provide information about lichen frequency, common substrates, site characteristics and provincial ranges (see “Lichen Distribution in British Columbia,” below). The following terms and schema are adopted: (Rare, infrequent, frequent, or common) over (acid, base-rich, mossy, or seasonally inundated) rock, (coniferous or deciduous) trees, (decaying) wood, moss, duff, or soil in (exposed, open, sheltered, or shady) (provincial range) (old growth) forests, steppe, depressions, or outcrops, at lower or higher elevations (throughout). Lichen Distribution in British Columbia: Lichen distribution is expressed according to the terms listed in the first two columns of Table 1 and in part mapped in Figure 2. Distribution is occasionally expressed in biogeoclimatic units, and these are listed in the third column of Table 1 and mapped in Figure 3. For further notes on the Biogeoclimatic System, see “Understanding Biogeoclimatic Zonation,” page 5. Species of widespread occurrence in the province are described as occurring “throughout.” The corresponding units of continentality in the fourth column are based on Conrad’s Index of Continentality (Conrad 1946). These are included to enable ecoclimatic comparisons with other portions of the world (for further details, see Goward and Ahti 1992).

TABLE 1. Distributional units and their definition General range Coast

Biogeoclimatic equivalenta

Conrad’s Index of Continentality

Hypermaritime

CWH wh and vh

Maritime – dry – wet – subalpine

CDF CWH (not wh and vh) MH

Intermontane – semi-arid – dry – moist – wet – subalpine

BG, PP IDF SBS, SBPS, MS ICH ESSF

Boreal

BBWS, SWB

Subalpine

Throughout

MH, ESSF

various

Alpine

Throughout

AT

various

Widespread

Throughout

Throughout

various

Inland

a

Life zone

40

INTRODUCTION

Figure 2 "Life Zones" of British Columbia*

BOREAL

55o

INTERMONTANE

HYPERMARITIME Scale 50

50o

0

50

MARITIME

100 150 200

kilometres

* see page 5

130o

120o

FIGURE 2. “Life zones” of British Columbia.

Total Range in the Northern Hemisphere: Total range in the northern hemisphere is expressed relative to western North America in the following distributional units: • western N Am • western N Am – eastern N Am • western N Am – western Eurasia • western N Am – eastern Eurasia • incompletely circumpolar (= any three of the above distributional units) • circumpolar North–south ranges in western North America are summarized for each species using the following geographic units: N to AK (Alaska) or YU (Yukon); and S to AZ (Arizona), CA (California), CO (Colorado), ID (Idaho), MT (Montana), MX (Mexico), NM (New Mexico), NV (Nevada), OR (Oregon), UT (Utah), WA (Washington), or WY (Wyoming). Species considered to have the northern or southern limits of their range within the province are denoted as N to BC and S to BC, respectively. Reactions: Only positive spot test reactions to commonly used chemical reagents are given. For further details see “Making Use of Lichen Chemistry,” page 13. Contents: Here a listing of dominant lichen substances is presented in alphabetical order. Substances given in parentheses (...) do not occur in all specimens. Notes: This section is reserved for details pertinent to the determination or treatment of the species — taxonomic and nomenclatural problems, notes on similar species, chemistry, and keys to varieties and subspecies. 4

INTRODUCTION

Understanding Biogeoclimatic Zonation British Columbia is a highly diverse province in which hundreds of ecosystems can be recognized. Maintaining these in the face of increasing pressure for resource development represents an enormous challenge — and involves, as a first step, classifying the province’s ecosystems in detail. In recent years, researchers with the B.C. Ministry of Forests have described medium-scale ecosystems according to the principles of biogeoclimatic ecosystem classification (Pojar et al. 1987). They have also arranged these ecosystems into a hierarchical system of biogeoclimatic zones, subzones, and variants. Collectively, the zones, subzones, and variants of the biogeoclimatic system are referred to as biogeoclimatic units. Each unit is characterized by a unique set of climatic variables, and supports — and is for practical purposes defined by — a unique vegetation. In biogeoclimatic ecosystem classification, the defining vegetation for each unit occurs on moderately well-drained sites. Such sites are said to be “zonal.” The most encompassing of the biogeoclimatic units is the biogeoclimatic zone. Fourteen biogeoclimatic zones are recognized for British Columbia and many of these are used here to describe lichen distribution. They are briefly characterized in Table 2 and mapped in Figure 3. For a more detailed summary, see Ecosystems of British Columbia (Meidinger and Pojar 1991). Lichen distribution may also be expressed using more generalized classification systems such as the “life zone system” (see Figure 2) and “general range system” adopted here. These systems are compared with their biogeoclimatic counterparts in Table 1. The comparison is made mostly at the zonal level, though two biogeoclimatic subzones have also been used: the Wet Hypermaritime (wh) and Very Wet Hypermaritime (vh) subzones of the Coastal Western Hemlock Zone (CWH). These subzones occur in the hypermaritime or outer coastal areas of British Columbia (see Figure 2). See Table 2 for the full names of other biogeoclimatic zones. TABLE 2. Summary information on the biogeoclimatic zones of British Columbia (Source: Lavender et al. 1990) Selected climatic characteristicsa Monthly temp. range

Zone

Code

Zonal vegetation

Zonal soils

Alpine Tundra

AT

Cassiope spp., Phyllodoce spp., Luetkea pectinata, Loiseleuria procumbens, Dryas spp., Salix spp., Silene acaulis, Poa spp., Festuca spp., Carex spp., Cetraria spp., Stereocaulon spp., Polytrichum piliferum

Regosols, Humic Regosols, Brunisols, Humo-Ferric Podzols

Boreal White and Black Spruce

BWBS

White spruce, lodgepole pine, black spruce, Rosa acicularis, Viburnum edule, Mertensia paniculata, Pyrola asarifolia, Cornus canadensis, Vaccinium vitis-idaea, Ptilium crista-castrensis, Pleurozium schreberi

Gray Luvisols, Dystric and Eutric Brunisols

-24.5–16.6

Agropyron spicatum, Artemisia tridentata, Artemisia frigida, Poa sandbergii, Koeleria macrantha, Festuca scabrella, Festuca idahoensis, Chrysothamnus nauseous

Brown and Dark Brown Chernozems

-10.8–22.4

Bunchgrass BG

5

-11.1–9.5

°days > 5°C 427

°days 5°C

°days 5°C

°days 5°C

°days